Nitrogen fixation and nitrogen gene abundance of METEOR cruises M77/3 and M77/4

Nitrogen fixation, the biological reduction of dinitrogen gas (N2) to ammonium (NH4+), is quantitatively the most important external source of new nitrogen (N) to the open ocean. Classically, the ecological niche of oceanic N2 fixers (diazotrophs) is ascribed to tropical oligotrophic surface waters, often depleted in fixed N, with a diazotrophic community dominated by cyanobacteria. Although this applies for large areas of the ocean, biogeochemical models and phylogenetic studies suggest that the oceanic diazotrophic niche may be much broader than previously considered, resulting in major implications for the global N-budget. Here, we report on the composition, distribution and abundance of nifH, the functional gene marker for N2 fixation. Our results show the presence of eight clades of diazotrophs in the oxygen minimum zone (OMZ) off Peru. Although proteobacterial clades dominated overall, two clusters affiliated to spirochaeta and archaea were identified. N2 fixation was detected within OMZ waters and was stimulated by the addition of organic carbon sources supporting the view that non-phototrophic diazotrophs were actively fixing dinitrogen. The observed co-occurrence of key functional genes for N2 fixation, nitrification, anammox and denitrification suggests that a close spatial coupling of N-input and N-loss processes exists in the OMZ off Peru. The wide distribution of diazotrophs throughout the water column adds to the emerging view that the habitat of marine diazotrophs can be extended to low oxygen/high nitrate areas. Furthermore, our statistical analysis suggests that NO2- and PO43- are the major factors affecting diazotrophic distribution throughout the OMZ. In view of the predicted increase in ocean deoxygenation resulting from global warming, our findings indicate that the importance of OMZs as niches for N2 fixation may increase in the futur

(Table 2, page 50) Toolik lake sediment chemistry (0-1 cm interval)

The sediments within Toolik Lake in arctic Alaska are characterized by extremely low rates of organic matter sedimentation and unusually high concentrations of iron and manganese. Pore water and solid phase measurements of iron, manganese, trace metals, carbon, nitrogen, phosphorus, and sulfur are consistent with the hypothesis that the reduction of organic matter by iron and manganese is the most important biogeochemical reaction within the sediment. Very low rates of dissolved oxygen consumption by the sediments result in an oxidizing environment at the sediment-water interface. This results in high retention of upwardly-diffusing iron and manganese and the formation of metal-enriched sediment. Phosphate in sediment pore waters is strongly adsorbed by the metal-enriched phases. Consequently, fluxes of phosphorus from the sediments to overlying waters are very small and contribute to the oligotrophic nature of the Toolik Lake aquatic system. Toolik Lake contains an unusual type of lacustrine sediment, and in many ways the sediments are similar to those found in oligotrophic oceanic environments.

(Table 1) Radiocarbon dating of a lake sediment core, southern West Greenland

The dominant processes determining biological structure in lakes at millennial timescales are complex. In this study, we used a multi-proxy approach to determine the relative importance of in-lake versus indirect processes on the Holocene development of an oligotrophic lake in SW Greenland (66.99°N, 50.97°W). A 14C and 210Pb-dated sediment core covering approximately 8500 years BP was analyzed for organic-inorganic carbon content, pigments, diatoms, chironomids, cladocerans, and stable isotopes (d13C, d18O). Relationships among the different proxies and a number of independent controlling variables (Holocene temperature, an isotope-inferred cooling period, and immigration of Betula nana into the catchment) were explored using redundancy analysis (RDA) independent of time. The main ecological trajectories in the lake biota were captured by ordination first axis sample scores (18-32% variance explained). The importance of the arrival of Betula (ca. 6500 years BP) into the catchment was indicated by a series of partial-constrained ordinations, uniquely explaining 12-17% of the variance in chironomids and up to 9% in pigments. Climate influences on lake biota were strongest during a short-lived cooling period (identified by altered stable isotopes) early in the development of the lake when all proxies changed rapidly, although only chironomids had a unique component (8% in a partial-RDA) explained by the cooling event. Holocene climate explained less variance than either catchment changes or biotic relationships. The sediment record at this site indicates the importance of catchment factors for lake development, the complexity of community trends even in relatively simple systems (invertebrates are the top predators in the lake) and the challenges of deriving palaeoclimate inferences from sediment records in low-Arctic freshwater lakes.

Size-partitioned phytoplankton carbon concentrations retrieved from ocean color data, links to data in netCDF format

Owing to their important roles in biogeochemical cycles, phytoplankton functional types (PFTs) have been the aim of an increasing number of ocean color algorithms. Yet, none of the existing methods are based on phytoplankton carbon (C) biomass, which is a fundamental biogeochemical and ecological variable and the "unit of accounting" in Earth system models. We present a novel bio-optical algorithm to retrieve size-partitioned phytoplankton carbon from ocean color satellite data. The algorithm is based on existing methods to estimate particle volume from a power-law particle size distribution (PSD). Volume is converted to carbon concentrations using a compilation of allometric relationships. We quantify absolute and fractional biomass in three PFTs based on size - picophytoplankton (0.5-2 µm in diameter), nanophytoplankton (2-20 µm) and microphytoplankton (20-50 µm). The mean spatial distributions of total phytoplankton C biomass and individual PFTs, derived from global SeaWiFS monthly ocean color data, are consistent with current understanding of oceanic ecosystems, i.e., oligotrophic regions are characterized by low biomass and dominance of picoplankton, whereas eutrophic regions have high biomass to which nanoplankton and microplankton contribute relatively larger fractions. Global climatological, spatially integrated phytoplankton carbon biomass standing stock estimates using our PSD-based approach yield - 0.25 Gt of C, consistent with analogous estimates from two other ocean color algorithms and several state-of-the-art Earth system models. Satisfactory in situ closure observed between PSD and POC measurements lends support to the theoretical basis of the PSD-based algorithm. Uncertainty budget analyses indicate that absolute carbon concentration uncertainties are driven by the PSD parameter No which determines particle number concentration to first order, while uncertainties in PFTs' fractional contributions to total C biomass are mostly due to the allometric coefficients. The C algorithm presented here, which is not empirically constrained a priori, partitions biomass in size classes and introduces improvement over the assumptions of the other approaches. However, the range of phytoplankton C biomass spatial variability globally is larger than estimated by any other models considered here, which suggests an empirical correction to the No parameter is needed, based on PSD validation statistics. These corrected absolute carbon biomass concentrations validate well against in situ POC observations.

Organic carbon flux and remineralization in surface sediments

Organic carbon fluxes through the sediment/water interface in the high-latitude North Atlantic were calculated from oxygen microprofiles. A wire-operated in situ oxygen bottom profiler was deployed, and oxygen profiles were also measured onboard (ex situ). Diffusive oxygen fluxes, obtained by fitting exponential functions to the oxygen profiles, were translated into organic carbon fluxes and organic carbon degradation rates. The mean Corg input to the abyssal plain sediments of the Norwegian and Greenland Seas was found to be 1.9 mg C/m**2/d. Typical values at the seasonally ice-covered East Greenland continental margin are between 1.3 and 10.9 mg C/m**2/d (mean 3.7 mg C/m**2/d), whereas fluxes on the East Greenland shelf are considerably higher, 9.1-22.5 mg C/m**2/d. On the Norwegian continental slope Corg fluxes of 3.3-13.9 mg C/m**2/d (mean 6.5 mg C/m**2/d) were found. Fluxes are considerably higher here compared to stations on the East Greenland slope at similar water depths. By repeated occupation of three sites off southern Norway in 1997 the temporal variability of diffusive O2 fluxes was found to be quite low. The seasonal signal of primary and export production from the upper water column appears to be strongly damped at the seafloor. Degradation rates of 0.004-1.1 mg C/cm**3/a at the sediment surface were calculated from the oxygen profiles. First-order degradation constants, obtained from Corg degradation rates and sediment organic carbon content, are in the range 0.03-0.6/a. Thus, the corresponding mean lifetime of organic carbon lies between 1.7 and 33.2 years, which also suggests that seasonal variations in Corg flux are small. The data presented here characterize the Norwegian and Greenland Seas as oligotrophic and relatively low organic carbon deep-sea environments.

Geochemical measurements of the Tasmanian Gateway from ODP Hole 189-1172A

Tectonic changes that produced a deep Tasmanian Gateway between Australia and Antarctica are widely invoked as the major mechanism for Antarctic cryosphere growth and Antarctic Circumpolar Current (ACC) development during the Eocene/Oligocene (E/O) transition (34-33 Ma). Ocean Drilling Program (ODP) Leg 189 recovered near-continuous marine sedimentary records across the E/O transition interval at four sites around Tasmania. These records are largely barren of calcareous microfossils but contain a rich record of siliceous- and organic-walled marine microfossils. In this study we integrate micropaleontological, sedimentological, geochemical, and paleomagnetic data from Site 1172 (East Tasman Plateau) to identify four distinct phases (A-D) in the E/O Tasmanian Gateway deepening that are correlative among ODP Leg 189 sites. Phase A, prior to 35.5 Ma: minor initial deepening characterized by a shallow marine prodeltaic setting with initial condensation episodes. Phase B, 35.5-33.5 Ma: increased deepening marked by the onset of major glauconitic deposition and inception of energetic bottom-water currents. Phase C, 33.5-30.2 Ma: further deepening to bathyal depths, with episodic erosion by increasingly energetic bottom-water currents. Phase D, <30.2 Ma: establishment of stable, open-ocean, warm-temperate, oligotrophic settings characterized by siliceous-carbonate ooze deposition. Our combined evidence indicates that this early Oligocene Tasmanian Gateway deepening initially produced an eastward flow of relatively warm surface waters from the Australo-Antarctic Gulf into the southwestern Pacific Ocean. This "proto-Leeuwin" current fundamentally differs from previous regional reconstructions of eastward flowing cool water (e.g., a "proto-ACC") during the early Oligocene and thereby represents an important new constraint for reconstructing regional- to global-scale dynamics for this major global change event.

(Table 1) Lipid biomarkers in surface sediments from the Gulf of Genoa

A series of molecular organic markers were determined in surface sediments from the Gulf of Genoa (Ligurian Sea) in order to evaluate their potential for palaeo-environmental reconstructions. Allochthonous input can be characterized by the distributions of n-C29 and n-C31 alkanes, n-C26 and n-C28 alkanols and branched glycerol dialkyl glycerol tetraethers (GDGTs), whose concentrations are generally highest near the river mouths. In the open basin however, terrestrial n-alkanes and n-alkanols may have an additional, aeolian source. Autochthonous input is represented by crenarchaeol and isoprenoid GDGTs. Their concentrations are highest in the open basin showing the preference of Thaumarchaeota for oligotrophic waters. Indications of a significant degradation of sterols and C37 alkenones exclude these lipids as reliable productivity proxies. Using terrestrial and aquatic lipids as end-members allows estimating the percentage of terrestrial organic matter between 20% and 58% in the coastal area decreasing to 1 to 30% in the deep basin. The spatial distribution of sea surface temperature (SST) estimates using the alkenone-based UK'37 index is very similar to the autumnal (November) mean satellite-based SST distribution. Conversely, TEXH86-derived SST estimates are close to winter SSTs in the coastal area and summer SSTs in the open basin. This pattern reflects presumably a shift in the main production of Thaumarchaeota from the coastal area in winter to the open basin in summer. This study represents a major prerequisite for the future application of lipid biomarkers on sediment cores from the Gulf of Genoa.

Planktic foraminifera analyses of surface sediments and satellite data in the Canary Islands region

The Canary Islands region occupies a key position with respect to biogeochemical cycles, with the zonal transition from oligotrophic to nutrient-rich waters and the contribution of Saharan dust to the particle flux. We present the distribution of geochemical proxies (TOC, carbonate, d15N, d13Corg, C/N-ratio) and micropaleontological parameters (diatoms, dinoflagellates, foraminifera, pteropods), in 80 surface-sediment samples in order to characterise the influence of coastally upwelled water on the domain of the subtropical gyre. Results of the surface-sediment analyses confirmed the high biomass gradient from the coast to the open ocean inferred from satellite data of surface chlorophyll or SST. The distribution of total dinoflagellate cysts, the planktic foraminifera species Globigerina bulloides, the diatom resting spore Chaetoceros spp., and TOC concentration coincided well with the areas of strong filament production off Cape Ghir and Cape Yubi. The warm-water planktic foraminifera Globigerinoides ruber (white), the diatom Nitzschia spp., and the d15N-values showed the opposite trend with high values in the open ocean. Factor analyses on the planktic foraminifera species distribution indicated three major assemblages in the Canary Islands region that represent the present surface-water conditions from the upwelling influenced region via a mixing area towards the subtropical gyre.

Global distributions of diazotrophs nitrogen fixation rates - Depth integrated values computed from a collection of source datasets - Contribution to the MAREDAT World Ocean Atlas of Plankton Functional Types

The MAREDAT atlas covers 11 types of plankton, ranging in size from bacteria to jellyfish. Together, these plankton groups determine the health and productivity of the global ocean and play a vital role in the global carbon cycle. Working within a uniform and consistent spatial and depth grid (map) of the global ocean, the researchers compiled thousands and tens of thousands of data points to identify regions of plankton abundance and scarcity as well as areas of data abundance and scarcity. At many of the grid points, the MAREDAT team accomplished the difficult conversion from abundance (numbers of organisms) to biomass (carbon mass of organisms). The MAREDAT atlas provides an unprecedented global data set for ecological and biochemical analysis and modeling as well as a clear mandate for compiling additional existing data and for focusing future data gathering efforts on key groups in key areas of the ocean. The present data set presents depth integrated values of diazotrophs nitrogen fixation rates, computed from a collection of source data sets.

Dinoflagellates of sediment core GeoB1523-1

Large numbers of calcareous dinoflagellate cysts and the vegetative calcareous coccoid species Thoracosphaera heimii are generally found in sediments underlying oligotrophic and/or stratified (sub)surface water environments. It is difficult to distinguish between the relative importance of these two environmental parameters on calcareous cyst and T. heimii distribution as they usually covary, but this information is essential if we want to apply cysts properly in the reconstruction of palaeoenvironments and past surface water hydrography. In the multi-proxy core GeoB 1523-1 from the Ceará Rise region in the western equatorial Atlantic Ocean (covering the past 155 ka), periods of greatest oligotrophy are not synchronous with periods of greatest stratification (Rühlemann et al., 1996, doi:10.1016/S0025-3227(96)00048-5; Mulitza et al., 1997, doi:10.1130/0091-7613(1997)025<0335:PFAROP>2.3.CO;2; 335-338; Mulitza et al., 1998, doi:10.1016/S0012-821X(98)00012-0), giving us the unique opportunity to differentiate between the effects of both parameters on cyst accumulation. The calcareous cyst record of the core reflects prominent increases in accumulation rate of nearly all observed species only during the nutrient-enriched but more stratified isotopic (sub)stages 5.5, 5.3, 5.1 and 1. In this respect, the distribution trends in the core are more similar to those of the eastern equatorial upwelling region (GeoB 1105-4) than they are to those of the oligotrophic north-eastern Brazilian continental slope (GeoB 2204-2), even though temporal changes in bioproductivity are principally in antiphase between the eastern and western equatorial regions. We conclude that stratification of the upper water column and the presence of a well-developed thermocline are probably the more important factors controlling cyst distribution in the equatorial Atlantic, whereas the state of oligotrophy secondarily influences cyst production within a well-stratified environment.

Benthic foraminifera in surface sediments, South Atlantic

Sixty surface sediment samples from the eastern South Atlantic Ocean including the Walvis Ridge, the Angola and Cape basins, and the Southwest African continental margin were analysed for their benthic foraminiferal content to unravel faunal distribution patterns and ecological preferences. Live (stained with Rose Bengal) and dead faunas were counted separately and then each grouped by Q-mode principal component analysis into seven principal faunal end-members. Then, multiple regression technique was used to correlate Recent assemblages with available environmental variables and to finally differentiate between four principal groups of environmental agents acting upon the generation of benthic foraminiferal assemblages: (1) seasonality of food supply and organic carbon flux rates, together with oxygen content in the pore and bottom waters; (2) lateral advection of deep-water masses; (3) bottom water carbonate corrosiveness; and (4) energetic state at the benthic boundary layer and grain size composition of the substrate. Food supply and corresponding dissolved oxygen contents in the pore and bottom waters turned out to be the most important factors which control the distribution pattern of the Recent benthic foraminifera. At the continental margin, in the zone of coastal upwelling and its mixing area, benthic foraminiferal assemblages are dominated by stenobathic high-productivity faunas, characterized by elevated standing stocks, low diversities and a large number of endobenthic living species. At the continental shelf and upper continental slope the live assemblages are characterized by Rectuvigerina cylindrica, Uvigerina peregrina s.1., Uvigerina auberiana and Rhizammina spp. while the dead assemblages are characterized by Cassidulina laevigata, Bolivina dilatata, Bulimina costata and B. mexicana. At the lower continental slope strong influence of high organic matter fluxes on the species composition is restricted to the area off the Cunene river mouth, where the live assemblage is dominated by Uvigerina peregrina s.1., the corresponding dead assemblage by Melonis barleeanum and M. zaandamae. In the adjacent areas of the lower continental slope the biocoenosis is characterized by Reophax bilocularis, and Epistominella exigua which becomes dominant in the corresponding dead assemblage. At the Walvis Ridge and in the abyssal Angola and Cape basins, where organic matter fluxes are low and highly seasonal, benthic foraminiferal assemblages reflect both the oligotrophic situation and the deep and bottom water mass configuration. The top and flanks of the Walvis Ridge are inhabited by the Rhizammina, Psammosphaera and R. bilocularis live assemblages, the corresponding dead assemblages are dominated by G. subglobosa on the ridge top and E. exigua on the flanks. Within the highly diverse E. exigua dead assemblage several associated epibenthic species coincide with the core of NADW between about 1600 and 3700 m water depth. These species include Osangularia culter, Cibicidoides kullenbergi, Melonis pompilioides, Bolivinita pseudothalmanni and Bulimina alazanensis. The assemblages of the abyssal Cape and Angola basins are characterized by Nuttallides umbonifer and a high proportion of agglutinated species. These species are adapted to very low organic matter fluxes and a carbonate corrosive environment.

Abundance of calcareous dinoflagellates of sediment cores from the South Atlantic Ocean

Despite the increasing interest in the South Atlantic Ocean as a key area of the heat exchange between the southern and the northern hemisphere, information about its palaeoceanographic conditions during transitions from glacial to interglacial stages, the so-called Terminations, are not well understood. Herein we attempt to increase this information by studying the calcareous dinoflagellate cysts and the shells of Thoracosphaera heimii (calcareous cysts) of five Late Quaternary South Atlantic Ocean cores. Extremely high accumulation rates of calcareous cysts at the Terminations might be due to a combined effect of increased cyst production and better preservation as result of calm, oligotrophic conditions in the upper water layers. Low relative abundance of Sphaerodinella albatrosiana compared with Sphaerodinella tuberosa in the Cape Basin may be the result of the relatively colder environmental conditions in this region compared with the equatorial Atlantic Ocean with high relative abundance of S. albatrosiana. Furthermore, the predominance of S. tuberosa during glacials and interglacials at the observed site of the western Atlantic Ocean reflects decreased salinity in the upper water layer.

Delta 13C measured on alkenones of surface sediment samples GeoB1008-6 to GeoB3603-1 from the South Atlantic

We have analyzed the stable carbon isotopic composition of the diunsaturated C37 alkenone in 29 surface sediments from the equatorial and South Atlantic Ocean. Our study area covers different oceanographic settings, including sediments from the major upwelling regions off South Africa, the equatorial upwelling, and the oligotrophic western South Atlantic. In order to examine the environmental influences on the sedimentary record the alkenone-based carbon isotopic fractionation (Ep) values were correlated with the overlying surface water concentrations of aqueous CO2 ([CO2(aq)]), phosphate, and nitrate. We found Ep positively correlated with 1/[CO2(aq)] and negatively correlated with [PO43-] and [NO3-]. However, the relationship between Ep and 1/[CO2(aq)] is opposite of what is expected from a [CO2(aq)] controlled, diffusive uptake model. Instead, our findings support the theory of Bidigare et al. (1997, doi:10.1029/96GB03939) that the isotopic fractionation in haptophytes is related to nutrient-limited growth rates. The relatively high variability of the Ep-[PO4] relationship in regions with low surface water nutrient concentrations indicates that here other environmental factors also affect the isotopic signal. These factors might be variations in other growth-limiting resources such as light intensity or micronutrient concentrations.

Distribution of diatoms, coccolithophores and planktic foraminifera in the Arabian Sea

The distribution of diatoms, coccolithophores and planktic foraminifers mirrored the hydrographic and trophic conditions of the surface ocean (0-100 m) across the upwelling area off the Oman coast to the central Arabian Sea during May/June 1997 and July/August 1995. The number of diatoms was increased in waters with local temperature minimum and enhanced nutrient concentration (nitrate, phosphate, silicate) caused by upwelling. Vegetative cells of Chaetoceros dominated the diatom assemblage in the coastal upwelling area. Towards the more nutrient depleted and stratified surface waters to the southeast, the number of diatoms decreased, coccolithophore and planktic foraminiferal numbers increased, and floral and faunal composition changed. In particular, the transition between the eutrophic upwelling region off Oman and the oligotrophic central Arabian Sea was marked by moderate nutrient concentration, and high coccolithophore and foraminifer numbers. Florisphaera profunda, previously often referred as a 'lower-photic-zone-species', was frequent in water depths as shallow as 20 m, and at high nutrient concentration up to 14 µmol NO3/l and 1.2 µmol PO4/. To the oligotrophic southeast of the divergence, cell densities of coccolithophores declined and Umbellosphaera irregularis prevailed throughout the water column down to 100 m depth. In general, total coccolithophore numbers were limited by nutrient threshold concentration, with low numbers (<10*10**3 cells/l) at high [NO3] and [PO4], and high numbers (>70*10**3 cells/l) at low [NO3] and [PO4]. The components of the complex microplankton succession, diatoms, coccoliths and planktic foraminifers (and possibly others), should ideally be used as a combined paleoceanographic proxy. Consequently, models on plankton ecology should be resolved at least for the seasonality, to account for the bias of paleoceanographic transfer calculations.

Living benthic foraminifers from the central Arctic Ocean

Fifty short sediment cores collected with a multiple corer and five box cores from the central Arctic Ocean were analysed to study the ecology and distribution of benthic foraminifers. To work out living faunal associations, standing stock and diversity, separate analyses of living (Rose Bengal stained) and dead foraminifers were carried out for the sediment surface. The size fractions between 63 and 125 µm and >125 µm were counted separately to allow comparison with former Arctic studies and with studies from the adjacent Norwegian-Greenland Sea, Barents Sea and the North Atlantic Ocean. Benthic foraminiferal associations are mainly controlled by the availability of food, and competition for food, while water mass characteristics, bottom current activity, substrate composition, and water depth are of minor importance. Off Spitsbergen in seasonally ice-free areas, high primary production rates are reflected by high standing stocks, high diversities, and foraminiferal associations (>125 µm) that are similar to those of the Norwegian-Greenland Sea. Generally, in seasonally ice-free areas standing stock and diversity increase with increasing food supply. In the central Arctic Ocean, the oligotrophic permanently ice-covered areas are dominated by epibenthic species. The limited food availability is reflected by very low standing stocks and low diversities. Most of these foraminiferal associations do not correspond to those of the Norwegian-Greenland Sea. The dominant associations include simple agglutinated species such as Sorosphaerae, Placopsilinellae, Komokiacea and Aschemonellae, as well as small calcareous species such as Stetsonia horvathi and Epistominella arctica. Those of the foraminiferal species that usually thrive under seasonally ice-free conditions in middle bathyal to lower bathyal water depth are found under permanently ice-covered conditions in water depths about 1000 m shallower, if present at all.